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Bothriolepid Antiarchs (Vertebrata, Placodermi) from the Devonian of the North-Western Part of the East European Platform

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Bothriolepid Antiarchs (Vertebrata, Placodermi) from the Devonian of the North-Western Part of the East European Platform Bothriolepid antiarchs (Vertebrata, Placodermi) from the Devonian of the north-western part of the East European Platform Erwin LUKŠEVIČS Institute of Geology, University of Latvia, Rainis Blvd 19 Rīga LV 1586 (Latvia) [email protected] [email protected] Lukševičs E. 2001. — Bothriolepid antiarchs (Vertebrata, Placodermi) from the Devonian of the north-western part of the East European Platform. Geodiversitas 23 (4) : 489-609. ABSTRACT A large collection of bothriolepid antiarch material from 26 localities in the north-western part of the East European Platform (Latvia, Lithuania, and the Leningrad, Novgorod, and Pskov regions of Russia) is described and illustrat- ed, with a critical treatment of previously described material. The genus Grossilepis Stensiö, 1948 is represented by two species, Grossilepis tuberculata (Gross, 1941) and G. spinosa (Gross, 1942). All remaining material is referred to the genus Bothriolepis Eichwald, 1840: B. prima Gross, 1942, B. obrutschewi Gross, 1942, B. cellulosa (Pander in Keyserling, 1846), B. pan- deri Lahusen, 1880, B. traudscholdi Jaekel, 1927, B. maxima Gross, 1933, B. evaldi Lyarskaja, 1986 come from the Givetian(?)-Frasnian deposits, and B. leptocheira curonica Gross, 1942, B. ornata Eichwald, 1840, B. jani Lukševičs, 1986, B. heckeri n. sp., B. ciecere Lyarskaja in Lyarskaja & Savvaitova, 1974 as well as one unnamed form come from the Famennian KEY WORDS sequence. The stratigraphic distribution of bothriolepids through the Vertebrata, Placodermi, Middle/Upper Devonian of the Main Devonian Field is clarified in a proposed antiarchs, new version of the vertebrate zonation. A cladistic analysis of the distribution Devonian, Baltic area, of several features within the family Bothriolepididae using the PHYLIP pro- biostratigraphy. gram package provides a new version of the interrelationships of bothriolepids. GEODIVERSITAS • 2001 • 23 (4) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.mnhn.fr/publication/ 489 Lukševičs E. RÉSUMÉ Bothriolépides antiarches (Vertebrata, Placodermi) du Dévonien du nord- ouest de la plateforme est-européenne. Une large collection de Bothriolepididae de 26 gisements du nord-ouest de la plateforme est-européenne (Lettonie, Lithuanie, régions de Leningrad, Novgorod et Pskov de la Russie) est étudiée et figurée, avec un traitement cri- tique du matériel précédemment décrit. Le genre Grossilepis Stensiö, 1948 est représenté par deux espèces : Grossilepis tuberculata (Gross, 1941) et G. spinosa (Gross, 1942). Le reste du matériel est attribué au genre Bothriolepis Eichwald, 1948 : B. prima Gross, 1942, B. obrutschewi Gross, 1942, B. cellulosa (Pander in Keyserling, 1846), B. panderi Lahusen, 1880, B. traudscholdi Jaekel, 1927, B. maxima Gross, 1933, B. evaldi Lyarskaja, 1986 du Givétien (?)-Frasnien, et B. leptocheira curonica Gross, 1942, B. ornata Eichwald, 1840, B. jani Lukševičs, 1986, B. heckeri n. sp., B. ciecere Lyarskaja in Lyarskaja & Savvaitova, 1974, et MOTS CLÉS Vertebrata, une forme non nommée d’âge famennien. La répartition stratigraphique des Placodermi, Bothriolepididae du Dévonien moyen et supérieur du Champ Dévonien antiarches, Principal et une nouvelle division en zones ichthyologiques sont proposées. Une Dévonien, Baltique, analyse cladistique des Bothriolepidae utilisant le logiciel PHYLIP a déjà fourni biostratigraphie. une nouveau scheme de relations de parenté des bothriolépides. CONTENTS Introduction ..................................................................................................................................... 491 Technique and terminology ............................................................................................................ 491 Historical account ............................................................................................................................ 494 Distribution and biostratigraphy .................................................................................................... 497 Lithologies, facies variations and vertebrate assemblages ............................................................ 499 Systematics ........................................................................................................................................ 505 Bothriolepis prima Gross, 1942 ....................................................................................................... 505 Bothriolepis obrutschewi Gross, 1942 ............................................................................................. 510 Bothriolepis cellulosa (Pander in Keyserling, 1846) ....................................................................... 517 Bothriolepis panderi Lahusen, 1880 ................................................................................................ 519 Bothriolepis traudscholdi Jaekel, 1927 ............................................................................................ 521 Bothriolepis maxima Gross, 1933 ................................................................................................... 534 Bothriolepis evaldi Lyarskaja, 1986 ................................................................................................ 539 Bothriolepis leptocheira Traquair, 1893 .......................................................................................... 544 Bothriolepis ornata Eichwald, 1840 ................................................................................................ 554 Bothriolepis jani Lukševičs, 1986 .................................................................................................... 569 Bothriolepis heckeri n. sp. ................................................................................................................ 577 Bothriolepis ciecere Lyarskaja in Lyarskaja & Savvaitova, 1974 .................................................. 578 Bothriolepis sp. indet. 1 .................................................................................................................... 594 Grossilepis Stensiö, 1948 .................................................................................................................. 595 Grossilepis tuberculata (Gross, 1941) ............................................................................................. 595 Grossilepis spinosa (Gross, 1942) ..................................................................................................... 598 Interrelationships of Baltic bothriolepids ...................................................................................... 598 Acknowledgements .......................................................................................................................... 603 References ......................................................................................................................................... 604 Appendix............................................................................................................................................ 609 490 GEODIVERSITAS • 2001 • 23 (4) Bothriolepids (Vertebrata, Placodermi) from the Devonian of Baltic INTRODUCTION describe and illustrate the armour structure of each species as completely as possible, because The family Bothriolepididae Cope, 1886 is the the members of the genus Bothriolepis are con- most diverse group of antiarchs in the western siderably variable in the shape of individual part of the East European Platform, represented plates and their overlap patterns as well as in the by at least 15 taxa. Subdivision of the Upper course of sensory line canals. Devonian of the Baltic region and north-west- The largest part of studied and described speci- ern part of Russia is based largely on fish data. mens was collected by J. Blesse, L. Lyarskaja, Bothriolepididae are very important for the cor- W. Gross, N. Delle, V. Sorokin, J. Upenieks and relation of Frasnian and Famennian deposits of the author, and is housed in the Latvian the Main Devonian Field of the East-European Museum of Natural History (Rīga). Other stud- Platform with similar strata elsewhere. The ied material is in the Paleontological Institute, genus Bothriolepis is of particular interest as it is Moscow (coll. D. Obruchev, R. Hecker, found in the Middle and Late Devonian deposits J. Eglons, N. Krupina, F. Chernyshov); Institute on all continents. of Geology of Lithuania, Vilnius (coll. Gross (1933, 1941, 1942), Lyarskaja (1978, 1981, V. Talimaa); Mining Museum, Saint-Petersburg 1986), Obruchev (1928, 1947, 1958, 1964), (coll. H. Helmersen, I. Lahusen, A. Olivieri); Karatajūte-Talimaa (1963, 1966) and others pro- Institute of the Earth Crust, St.-Petersburg (coll. vided lists of agnathans and fishes characteristic A. Ivanov, W. Ermolaew, S. Snigirevsky); for various Devonian beds in the Baltic and Museum of Geology of the University of Latvia, north-western part of Russia, described verte- Rīga (coll. N. Delle, R. Kampe); Natural brate assemblages and correlated some strata History Museum, London (coll. P. Egerton, with the Upper Devonian of other regions. Six J. Eglons, R. Gross); and the National Museum new species have been described from the Main of Scotland, Edinburgh (coll. R. Gross, Devonian Field since Stensiö’s (1948) major H. Trautschold, J. V. Rohon). revision of the Bothriolepididae. During prepa- ration of this work, a large amount of bothriole- pidid remains stored in the main museums of TECHNIQUE AND TERMINOLOGY Latvia, Lithuania, Russia and Great Britain was studied. A large
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